Don never had a phone at home, he never used a computer or had an email account, he walked to work, he foraged for food in the urban landscape--harvesting watercress from Ravenna Creek and collecting fennel and mushrooms from where he planted them in his walks around the city.

At work, Don could not bring himself to throw anything away. I always knew that, but it became even more obvious to me as I began to look through his desk and bench in the lab. For those of you who came by Don's area of the lab, I am sure you must have noticed the mounds of mail and papers cascading from his desk and onto his lab bench. I don't know why he saved old paper towels with notes scrawled by lab mates telling him that his dentist's office had called or why he saved cut-up, unused checkbooks for accounts that were no longer active, but he did.

In the lab he rarely used a clean piece of paper, preferring instead to use the junk mail he received as note paper to work out the details of his experiments. I found a packet of pre-cut pieces of scrap paper ready for his future experiments. Some of the really important notes made it into notebooks, but to record the results of his crosses was the only time he used clean pieces of paper for his work.

When he did throw things away it was always done neatly, ceremoniously, almost reverentially. Old used plastic petri dishes were carefully stacked into a paper grocery bag. Only when the bag was complete full did he neatly fold down the top, staple it shut, and then carefully place it in the trash. Among the things I found on his bench was a piece of junk mail that had become the final resting place for bits of tape that could have no future use.

It was clear that he also foraged among the trash bins in the department. As I unearthed an old volt meter in the depths of one of his desk drawers, I recognized it immediately as something we threw out years ago because of the worn case and the cracked dial. I'm not even sure that it still worked. But Don quietly retrieved it.

Don was the master of reusing materials: toothpicks and tongue depressors are smooth and brown from decades of use. Of course his velvets, of which he had over 200, showed the wear of decades, also. In a way I think Don would be happy to know that his velvets have been recycled to another purpose. A quilt, I am sure, would never have occurred to Don, but I hope he would be pleased that some of his velvets will see future use in this new form. It seems fitting that these velvets be passed to his grand niece, Megan Harvey, who is following in her great uncle's footsteps as a budding microbiologist.

I really enjoyed knowing Don for over 30 years. I guess I really didn’t notice his presence in the Genetics Department until I got interested in yeast a couple of years after coming to Seattle and began attending the weekly yeast meetings. Those meetings had already been going on for several years, and Don seemed to have been a constant participant extending back to the time when he was a graduate student with Howard Douglas in Microbiology. Repeatedly, I was amazed at the depth of Don’s understanding of complex issues in yeast genetics then, and I never had any reason to think otherwise over the next 3 decades.

Don shared a lab with Herschel Roman, and the two of them were very different sorts. While Hersch was a commanding presence in every occasion, Don was retiring in his manner and cryptic in nearly everything I ever heard him say. Herschel rarely did experiments, but would beat the drums for yeast as the best experimental organism of the age (which turned out to be true), while Don simply did his own experiments elegantly and seemed to revel in the complexity of the results. Don was famous locally, of course, for being sort of helpful to the uninitiated, but gleeful in forcing them to think for themselves. Many told tales of his having given them strains that corresponded precisely with what they said they wanted, as opposed to what he knew they really wanted – so they got triploids, trisomics, strains with suppressors in them, and all sorts of other devious tests of the naïve recipient’s ability to recognize aberrant genetic behavior and solve a real puzzle. I got many strains from Don, and I don’t think he ever pulled this on me – but maybe he did and I never figured it out!

I know that Hersch got a bit frustrated with Don over the years for his reluctance to toot his own horn. Don had attacked and solved some very knotty problems, but didn’t broadcast the results very widely in print. I know Don had made fundamental contributions to some of the more significant advances in yeast genetics in that era – including nonsense suppression, mating type control, and galactose regulation. But his name was not widely associated with these findings because, while others were building on these foundations, Don had turned his attention to other problems rather than milking his successes.

In later years, Don became interested in meiotic recombination in hybrids between distantly related species. His favorite species for this work included one he’d isolated from spoiled mayonnaise. He’d originally decided it was a new species and designated it S. douglasii in honor of his thesis mentor, but Don later reluctantly decided that it really was an isolate of a described species, S. paradoxus. Another notable isolate came from spoiled orange juice and became known as OJ, until Mr. Simpson’s reputation plummeted. Hybrids between these strains and Baker’s yeast occupied much of Don’s thinking and experimentation for several years, especially during his many sojourns to Gif and other labs near Paris, where he also worked on other issues that he rarely mentioned to me. He was more forthcoming in discussing which French restaurants he had visited and which fungi he had managed to collect. When pressed about fine dining, however, it usually was revealed that he’d stayed at the lab most of his time there and eaten in the cafeteria.

Don lived simply in an apartment within walking distance of the lab. When I first got to know him, he would go to Lake City in the evening and have dinner at an inexpensive all-you-can-eat cafeteria called the Royal Fork. Later, he began having dinner more often (and even more frugally) at the McDonalds one block from his apartment. He could afford to live a little better than this, of course, but he seemed not to see any point in it. Obviously, his very generous bequest to our department is of such magnificent proportions partly because he lived so simply.

One of Don’s few extravagances was his cherished Triumph, which I think he’d bought just as soon as he could afford it. But he wasn’t one to flaunt his wealth, and the Triumph spent most of its time in the garage while Don walked to work. When the Department of Genetics began having departmental retreats at Friday Harbor in about 1975, these were the perfect occasions for Don to get the Triumph out of the garage and give someone a ride to or from the meetings. I’m not sure how much else he used the car, but I know that once he drove it all the way to a yeast conference in Banff and then offered me a ride home. I was glad to accept, and we spent two days driving all the way back to Seattle by way of Jasper and central British Columbia with the top down. I’ll never forget it.

Don was not only a very good scientist but also a very good person. I am very sad because I have lost a very good friend. Since the first time we began to work together in Gif, many years ago, we had a lot of interesting talks. Together we have gone to several scientific meetings held in Europe (traveling by plane, by train or with my car) and outside of Europe. The last time Don came to Paris and Gif, I drove him with my car going through different streets and spots in Paris. I am glad he could see those spots a last time a few weeks before dying because Don had a very good knowledge about Paris and liked very much this town. I have not only lost an excellent scientist colleague but also a really great friend. I am very sad.

A second note from Maurice:

I met Don hawthorne in 1969. In this period I worked for my thesis in the Center of Molecular Genetics of the CNRS in Gif-sur-Yvette. During Don's stay in Gif I worked with him on yeast strains carrying amber suppressor. We rapidly became friends. I appreciated Don's great knowledge and modesty. After this period Don came back often to Gif and each time we worked together. We took part together in many congresses in Europe and outside. During the last years, although his health was declining, Don continued coming twice a year to Gif to stay in my laboratory. We had a lot of interesting talks on yeast science and other purposes, generally in English but sometimes also in French because Don could understand and speak French. For this reason I finish this letter with French sentences to say:

Adieu Don, tu as bien rempli ta vie et tu laisses une oeuvre scientifique marquante. La dernière fois que tu es venu à Gif, en décembre 2002, tu pensais encore à de nouvelles expériences et à de nouvelles constructions de souches. L'annonce de ta mort m'a rempli d'une infinie tristesse et ton souvenir ne me quitte pas. Merci pour ton savoir, ta modestie, ta gentillesse et ton amitié.

Adieu Don.
Maurice

I'm really saddened to hear of Don's death. I know that he has been missed in the lab and his loss will be felt greatly. He contributed so much to yeast genetics and to the UW Department of Genetics/Genome Sciences over the years, and in particular to the atmosphere in the Fangman-Brewer lab. I always enjoyed the sense of history and timelessness that his presence lent the lab, and the fact that he was resolutely unimpressed by the hassles and vagaries that can obscure the real interest of science. He had quite a sense of humor too--I will never forget his driving me up the I-5 to Anacortes in his convertible that first September I was in Seattle, on the way to Friday Harbor. Having the chance to re-acquaint myself with Don was one of the many pleasures of my recent visit to Seattle. He is someone I am very glad to have had the opportunity to know.

Don was a special person. Extraordinarily intelligent and kind; a gentle, sensitive man with a wry sense of humor that revealed itself in quiet, playful ways in private and public forums. When I was a graduate student in Seattle, I occupied a lab next door to his. In the beginning I was quite shy and hardly talked to him. But towards the end of my first year that changed. My roommate and I invited him to dinner one night and he showed up with two bottles of wine and charmingly regaled us with stories of his travels and adventures around the world. It kind of broke the ice in my relationship with him, and from then on he often came into the lab to tell me stories…..about walking on old Roman roads in France and Italy, great restaurants I should go to, particularly in France, and about the work he was doing. Although on the surface he appeared to be a very reserved person, I came to understand that he really enjoyed people and delighted in conversation, especially with students and postdoctorals. And he was a great tease!

He loved to test our powers of deduction by presenting his work in the form of a puzzle, as sort of a little detective story. He would then watch keenly and with bemused interest, (I always thought secretly cheering us on), as we racked our brains trying to solve the mystery! I remember one evening when he stopped by to show me a greenish-blue galactose plate with lots of different yeast patches. They were varying shades of yellow, orange and blues due to a dye in the medium, which changed color with pH, as the galactose was utilized. Don quietly placed the plate on the table in from of me and looking straight at me with his clear blue eyes showing a hint of amusement in them said, "I just got these results from an experiment I did…..guess what the experiment was!"

And, one time, at a research presentation, he boldly challenged the whole department all at once. He said, "Uh, I have a problem for you all to solve… I have a yeast strain with these four markers and I found a mutation that segregates as a single gene which causes them to disappear and three new ones to appear." Glancing around the room, with an air of mischievousness, he said "Guess what the mutation is?" Silence followed. Don thrived on it! He patiently waited, dismissing one suggestion after another. Finally, after about a half hour, to his surprise and genuine pleasure, one student correctly figured it out shouting out: "I got it, I got it! It’s a tRNA mutation that changes an ochre to an amber suppressor!" "That’s right" said Don, "congratulations!"

Over the years, after leaving Seattle, when I returned to visit he would invite me to lunch or dinner and tell me stories of his early days in yeast genetics which I loved to hear. One evening, we went out to Ray’s Boathouse. During dinner, he spent about 3 hours telling me about the first yeast ascus he ever dissected, everything that was happening in the world at the time, and everything he did over the next several days while the spores grew up! And then came the exciting moment when he discovered that the spores wouldn’t mate, but they could sporulate and his realization that the original cell must have been a tetraploid! More than 30 years later, he obviously still took great delight in figuring it all out! I had a tremendous urge to run over to the dinner table next to ours and say…"Listen, did you hear that story! Wasn’t that great?"

Don loved science. It was an adventure. I believe that he regarded life the same way. He was a genuinely fine man and we will all miss him dearly.

I saw Don last year during a short visit in your lab, and he showed me plates where cells with very peculiar phenotypes were growing, exactly as he did 25 years ago. It was so nice to see that he still had the possibility to do genetics.

In 1975, I came from Paris, as a post-doc scientist in Herschel Roman’s laboratory. Don Hawthorne was there and it turned out to be a great chance for me. I knew his important contributions in yeast genetics, and was quite fascinated by his research. I learnt a lot from him, notably how to approach a genetic question and how to deal with it. Besides a common interest in genetics, we progressively discovered other common tastes, and a good souvenir is some beautiful escapades for picking mushrooms, together with Glenn Kawasaki, a bright young geneticist also working in Herschel’s lab.

After that time, I saw Don from times to times in France. He came almost every year to work in Gif-sur-Yvette with Piotr Slonimski. He was also exploring France, specially the Sud-Ouest, a very attractive area where food is indeed very good. Once, he came to our house with an unbelievable bottle of Cognac 1942 that we started to drink with great respect. It was also so interesting to see the roman coins that he collected and to listen to the story of the roman emperors, story that he knew quite well. It was always a great pleasure to see him. Now, he will remain in my mind and heart.

Don was a unique person in so many ways, and his absence brings about yet another end of an era. But I count myself fortunate to have known him, his sparkle, his quiet and sometimes mischievous ways. How sad to think of his corner in the lab empty.

My guess is that Don Hawthorne found me interesting. We first met at a Regulation Gordon Conference in the early '70s, where Don gave one of the featured talks. His presentation mystified the audience. Don's sparse lecturing style demanded intense concentration and a bent for cryptography. He delivered the information without affect or context, boiled down to the clues that had led him to his conclusion--which he revealed at the finale with what I interpreted as a satisfied half-smile. The enormous importance of the now eponymous "Hawthorne deletion", which was the focus of the talk, escaped the notice of an audience unfamiliar with the intrigues of yeast genetics. At the end of his talk, I approached him with a question. He responded with a curt, "You understood some of it but you didn't get it all." I felt somewhat shunned by this acerbic answer. The next morning at breakfast, Don sat down next to me and, without interruption, explained at length how he had arrived at his model. I interpreted his unexpected sociability not as remorse, but rather recognition of my potential aptitude for "getting it." I learned to enjoy this Sherlock Holmes/Dr. Watson relationship.

Don was truly a geneticist's geneticist. He absolutely reveled in the beauty of genetic complexity and was rather openly disdainful of the genetically unsophisticated. I remember him once almost cackling over the possibility of sending some unsuspecting molecular biologist the stock they had asked for (which they clearly did not understand) versus the stock they really wanted. Don thrilled in saying that the requested stock contained a bit of "a fun genetic puzzle" (as I recall, either a disomy or a chromosome aberration of some kind), one that might keep these poor molecular biologists "busy for a good little while". But in the end, he mailed them not what they asked for, but what they needed.

Don did genetics just because he loved it. Some fifteen years ago he showed me the results of introgressing a carlbergensis chromosome into a standard cerevisiae background. He demonstrated nicely that homeologs segregated from one another but did not recombine. I urged him to publish this result, but he demurred telling me that this experiment was "just a bit of genetic trickery" that he did solely for the joy of doing genetics.

The little dapper man with the bow-tie just really loved doing genetics. He did science for the pure pleasure of thinking genetically. I fear that breed grows far closer to extinction with Don's passing.

There are so many stories.

My favorite concerns my trying to do spore-cell matings under Don's tutelage using his micromanipulator. I struggled and struggled without success. Looking over my shoulder occasionally, Don finally suggested that I take a lunch break and come back "fresh". I took this advice and was very surprised that in my absence he took pity on me and completed the micromanipulation. I happily followed the fate of this experiment, taking photographs over the next day or so. When I wanted to remove the film and develop it Don was aghast because I hadn't used every frame. I explained that I would pay for developing and replace his film with a new role, but Don replied that "I was missing the point"; I was wasting the remaining frames. "OK" I said, he could finish the roll, I would still pay to develop it. Over the next several weeks I frequently stopped by to ask if he had finished the roll, but he had not. About 6 months later Don found me in the medical school library to anxiously hand me my prints. He had searched for me for hours because he knew I was "in a hurry" to have the pictures. Unfortunately, so much time had past that I had completely forgotten about this experiment; I had not a clue what he was showing me. The story emphasizes Don's kindness and generosity, superseded only by his frugality.

Another story: Jim and I sought Don's advice about traveling in France. We invited him to dinner to discuss places to see and people to meet. He came with 6 carrousels of slides and several bottles of French wine. I drank too much wine and missed the slide show. Don provided maps, books, suggested restaurants - all which turned out to be amazingly useful. Unfortunately we were robbed during our trip and we subsequently frantically searched bookstores, map stores, etc. for replacements of the items Don had lent to us. I can still see the hurt in his face when he realized that we had spent our postdoctoral salary funds to purchase these replacements; he never would have allowed us to do this if we had asked. Again amazing generosity, coupled with frugality.

Don was an amazing person: supersmart, highly cultured, generous, mischievous, frugal, and often cryptic. I miss him.

I remember Don Hawthorne with great respect and fondness. He taught a cytogenetics lab that I took as an undergraduate. It was a small class. We made preparations of chromosomes from various roots (onions, hyacinths--beautiful) and meiotic chromosomes from Tradescantia stamens. We saw fish chromosomes as well. He always kept us well supplied but was also willing to listen and allow students to follow their passions.

As an undergraduate I washed the dishes and made and poured the plates for Herschel Roman. Don made his own media for quite a while. Eventually, however, he let me make his. I felt quite honored to receive that degree of trust. I always admired Don's frugality as well. He simply never wasted media. Sydney Kustu once had a Polish postdoc who had worked in a lab in Poland with very limited resources. Sydney said that this postdoc could streak a hundred isolates for single colonies on a single Petri plate. Don would have understood and appreciated that postdoc. They had similar views on frugality and on the privilege it is to be able to do research, a privilege that one should not abuse.

I remember him well for the minor tests he would give to students when he thought they had progressed far enough to solve them. Receiving such a puzzle was viewed by us as a great honor, for it meant that one was progressing. I may have been the first student he gave such a puzzle to, for mine was pretty easy. I asked for a can1 strain. He streaked it out on YEPD and gave it to me. It also contained an arg mutation and hence wouldn't grow on synthetic media because it couldn't transport in the needed arginine. I discovered the puzzle after I crossed the strain and obtained nongrowing segregants (and parent, although it is not a certainty that I had that conrol on the plate). I figured it out fairly easily. Mike Esposito received a strain that had a translocation in it. That took longer to figure out.

Don was a modest man but one who knew his own worth. He made very substantial contributions to yeast genetics, contributions of which anyone would be proud. All of the gene mapping that provided the framework for others to build upon, the linear ascus and overlapping spindle story, nonsense suppressors, the galactose and melibiose tales, mating type switching, and so much else.

I owe Don a large debt of gratitude. He became my interim thesis advisor when Herschel Roman had a stroke. Don continued to mentor me when Herschel returned. Don and I played almost-daily games of 20 Questions on research results (for three years). I would visit his lab and would be shown yeast experiments. Then I had 20 questions to figure out the significance of the experiment. These sessions with Don were very challenging, and I learned a great deal of genetics from him.

We would continue our fungal experiments by visiting the Olympia Brewery twice a year and go mushroom hunting for chantrelles, boletes, and pseudomorels. After the trips in the woods, we'd go to Don's favorite eating place, The Royal Fork, and pig out. Don was an expert at buffet dining. For many years in Seattle, he would walk to the Royal Fork in the Wallingford neighborhood and would bring back his chicken bones to feed crows near his apartment in the U Village area. Don had many pets and dependents; they all had black feathers. During my graduate days, I invited Don to my place for Thanksgivings. He would be sure to take dinner scraps for his crows.

Eventually I got my Ph.D. and went to Boston. I returned to Seattle to become the first scientist at ZymoGenetics. Don and I, with Samir Deeb, continued to go on mushroom hunts, occasionally with my dog. In the mid-80s, Don and I attended conferences in Montpelier (France) and Rome (Italy) and traveled together to Milan, Venice, Paris, Monte Blanc, and other spots by Eurail. He knew all of the cheapest places to stay, usually near the train stations. It was a grueling four-weeks in Europe. We saw lots of churches and museums and ate well.

Over the years, Don and I saw less and less of each other. Occasionally, I would find him at the Genetics Department or run into him at a Presidents' Club function at the University (we were each Life Members). Or run into him at the QFC in the U Village. At the hospital, I thanked him for being my mentor and long-time friend. He was a brilliant scientist and a very generous person. It was an honor to know Don.

I worked in the lab across the hallway from Don for almost two years, that is from February 1974 through December 1975. When I came to the Department, Herschel explained to me that Don did not apply for research grants himself, but he was such a valuable geneticist that Herschel just wanted to have him in his lab.

Don had a large collection of yeast strains. Other people would have their strains individually on agar slant in the fridge. Not Don. He stabbed strains onto Petri plates supplemented with starch. He then let the plates dry down; the starch prevented the agar from cracking. I remember that Don often had many Petri plates on his bench with the lid off the dish and with paper towel covering the dish.

Don was a hard-core geneticist. If you wanted a strain in which some unusual genetic event had happened, like duplication of a chromosome or an illegitimate recombination, you would go to Don. Someone, I forget who, told me that he would sometimes give you a strain with an extra character that he did not tell you about and then come back and see if you had found out. I did not experience that myself, or perhaps I wasn't clever enough to find out.

I several times experienced Don giving an informal seminar for his yeast colleagues in the J-wing. His seminars were usually not very comprehensible. I think Don put an honor into keeping his seminar informal to the extent that he did not want to prepare his talk. When he talked about translational suppressors, e.g. nonsense suppressors, he liked to test his listeners for whether they could remember the genetic code by heart.

I remember Don being at work a lot, also in the weekends. Herschel had a technician, Julia (Richards). I talked with her about Don, whom she appreciated very much. She found it curious how he would get into his nice, somewhat formal dress and drive his sports car to the lab in the weekend and work there, and then drive home again.

Don was a friendly person. He had a characteristic twinkle in his eye together with his greeting smile, or when he wanted to implicate a deeper meaning in something he said. He did not seem to get really mad even in cases of disaster. Julia told me about an event shortly after she had started in the lab. She had noticed all these culture tubes in the shaker incubator that seemed to sit there day after day with nothing happening; she did not realize that they were Don's experiments, and that he would renew them all the time. After some weeks, Julia decided to tidy up, and she took the tubes out and emptied them. When Don then asked around for the whereabouts of his cultures, she had to explain him her mistake. He said that if he could just get the tubes, there would in all likelihood be yeast left to grow up again.

I shared a room with Don in Walt Fangman & Bonny Brewer's lab from 1990 to 1995. At this point in his career Don was, of course, a legendary figure in the world of genetics. The purity of his involvement in his work was one way that Don made a big impression on me. Like most scientists, I have always tried to maintain a balance between work and the rest of my existence. On the other hand, Don, as far as I could tell, sought happiness by making his work his entire life. I never saw Don in a bad mood (except twice during his bouts with cancer). This strongly suggests to me that he really was perfectly content in life. Would that we could all experience such contentment!

Don's science was from the era before genetics had the molecular tools it has today. It was his toolbox of classical genetic methods that I found most valuable. In particular, he performed a cytoductive cross that, literally, initiated my independent scientific career. On the other hand, he was so reserved, even furtive, about his own work that I'd be hard-pressed to tell you exactly what genetic problem he was actually interested in solving during that period I was his lab mate.

I would have loved to get to know Don better, but my several attempts to get together with him outside of work were politely rebuffed. One exception was a ride I got from him in his Triumph back to Seattle from Anacortes after the Genetics Department annual retreat at Friday Harbor. With the top down! I sure enjoyed this and I believe he enjoyed my company and showing off his spiffy car.

This is not the season to be an American Francophile, but Don and I had that in common over several decades. When bumping into him in the hallways of the J wing, the usual mutual opening of the conversation was "Are you planning to go back this year?" This would invariably bring forth a twinkle in those sometimes deceptively sleepy eyes and would result in a lively conversation about the language, the countryside, the wine, the food, the sunshine, and French geneticists (Ephrussi, Slonimski, Monnot, Jacob, Gros, among other colorful characters). And he would, of course, light up when the subject of nonsense suppressors would come up in a conversation. But as we all know, Don was a very private and reserved soul. Such waters run deep and are not easy to sample.

I'm very sad about Don's death. In 1984 I spent some time in his laboratory in the Department of Genetics in Seattle and we met regularly at the International Yeast Meetings. He was an excellent geneticist, did all experiments himself and I was always impressed, how he combined travels in France with yeast experiments.

As is usually the case, the death of a friend brings back a flood of fond memories, and Don's death has served that purpose for me. Many times recently I have wondered how he was. We had heard that he'd been ill. I thought of trying to get in touch with him although that was never easy. As far as I knew, he had no phone, and I had never known his address, although I had visited his apartment at various times, always driven there by himself. I think that this expressed his desire for privacy and independence, perhaps similar to my own.

Don's and my association began in the 1950s soon after Herschell Roman told me about Don's work on genetic mapping. At that time he was doing a post doc with Sterling Emmerson at Cal Tech. following completion of his graduate work at the U of W with Howard Douglas.

I was working at U.C. Berkeley isolating mutants in Saccharomyces cerevisiae and had inadvertently found a number of linkages between pairs of genes, and between genes and centromeres, which I thought could complement Don's work. I soon arranged a trip to Cal Tech to see Don. This was the beginning of our lengthy collaboration and 50 year friendship maintained through many meetings at Seattle, Berkeley, Italy and elsewhere during which we pursued our scientific, as well, as recreational interests.

After Don's return to Seattle, he resumed using his DeFonbrune micromanipulator and I brought a micromanipulator from Berkeley which had been designed by my mentor, Cornelius Tobias. For the next several years, we continued dissecting asci, making crosses and analyzing linkage data which provided the basis for the first yeast map which was published in "Genetics" in 1960. It is my view that this paper really helped establish the field of yeast genetics.

fermentation, and Lindegren whose work on neurospora he had transferred to his new interest in yeast, were pioneers in the field. Winge's work and publications were so complex that few people understood them. Lindegren's early work on neurospora was sound. But in his work on yeast, he tried to show that yeast didn't obey the rules of genetics--a mistaken idea! The field was ripe for our 1960 publication which showed that yeast obeyed the basic rules of genetics and thereby established a firm basis for the explosion of yeast genetics which soon followed.

Over the following ten years, we continued expanding the map, eventually publishing three more editions and a review of yeast genetics. At this point Don's interest turned elsewhere and I continued with other collaborators through edition XII.

Don's extremely careful and thorough lab techniques and his ability to hand-write almost final text facilitated a smooth and enjoyable collaboration.

However, our association was not all science. During my many trips to Seattle, he was a gracious host. I enjoyed touring the area in his various Triumphs, mushroom hunting, several fly fishing trips, visits to the family home in Olympia, and meeting some of his family. Not only did our work complement each other's but our personalities and recreational interests followed similar lines. We had many hours of quiet, comfortable companionship, whether in the laboratory or tramping the foggy, soggy undergrowth of the coastal hills north of San Fransisco hunting for mushrooms. One of our most exciting finds was a large Amanita clyptoderma, supposed to be edible, but being geneticists we were worried that it might have been a product of a cross between the edible and a deadly poisonous species. We knew of a fireman in the next town, who was reputed to be a mushroom authority, so we drove there. As we entered the fire station, he came toward us and upon seeing our mushroom, even from afar, he pointed to it and exclaimed in a large voice "That's a good one!" We took it back to our family cabin, ate it for supper and survived!

During his many trips to Berkeley, we also enjoyed tours of the wine country and Monterey area. One of our last adventures was touring the countryside and hill towns in Tuscany in his newest Triumph which he had driven down from Gif-sur-Yvette to Florence, where I was doing research on wine yeast at the Department of Genetics.

This reminds me of a little story of a later trip that he made to Florence by train. I went to the station to meet him at the appointed track and time. After everyone had exited the train, I waited for several minutes, then returned to my apartment wondering what had happened to him. The next day, he appeared at the Genetics Department at the University of Florence. He had been on the train, but apparently was in no hurry to get off. When he didn't find me at the station he took a taxi to the apartment, but couldn't find the apartment number, so went to a nearby hotel. This was typical of Don.

I have been so sorry to hear about Don. His contribution to the field of yeast genetics has obviously been a major one, even if sometimes I found hard to unravel his logic. It is true that he was a mysterious and intellectually attractive human being.

I remember when I arrived for the first time in Seattle in 1965 in order to spend a one year post-doc with Herschel Roman. Don (probably pushed by Herschel who was worried about this lonely young women just landing from Paris) invited me very formally to take me out for dinner. I was pleased and dressed in European standards for this occasion. At the planned day Don came to pick me up at 6 pm. We crossed the road and he announced that he was taking me to "the best hamburger place in town". Indeed the hamburger was delicious, the atmosphere was typically North Western of the time, solid wooden common tables and benches, juke boxes in the corners, young guys wearing blue jeans and boots, etc. During dinner, Don explained to me which sophisticated yeast crosses he was planning to perform. It is clear that although quite pleasant, the whole situation was different from good "old European" expectation. Among other experiences, this introduction with Don Hawthorne initiated my faithful love story with the Pacific North West and its peoples.

We send you a photo of Don taken in Seattle in 1985. This shows the Don very few people knew. We drove together with his sportscar through Seattle when I visited him during our ten years collaboration on S. douglasii and S. uvarum, two species that are probably very close to S. bayanus and S. paradoxus. He visited my lab regularly after his winter sabbaticals in France and each year we wrote a few more pages. This paper on speciation in Yeast may be one of the last publications of Don. I still have two reprints. The list of references in that paper contains other publications about S. douglasii with Don as co-author. By the way, it was submitted to Reed Wickner. When we did not receive reviews after two months I called him and he informed me that it was already in press. This was the only time I had submitted a publication which was written over a period of 8 years.

What do I remember most about Don? His passion for genetics, his passion for recycling, and his sly and somehat subversive sense of humor. I remember a classic Don moment at a Seattle Yeast meeting. Don had just given a talk (using recycled overhead transparencies on which he had re-traced the original faded writing) in which he had outlined a typically complicated genetic scheme to identify some gene. In the Q-and-A period, someone asked him why he didn't simply clone the gene. Don looked at him for a couple of seconds and responded, "Well, where's the mystique in that?"

Don had his finger on the pulse of popular culture--sort of. One day soon after it came out that the band Milli Vanilli hadn't really sung any of the songs they had supposedly done, Don stunned us by announcing that he was no longer going to purchase any of their albums. It was such a startling moment, it didn't really matter that he referred to the band as "Manilli Vanilli".

For years, Don had a low-grade running war with the campus groundskeepers. He would scatter fennel seeds outside Bonny's office, the groundskeepers would cut down the fennel plants, and the cycle would repeat. I never figured out if he did it because he particularly liked fennel or if it was just to have fun with the groundskeepers. For the last few months he seemed to have had the upper hand, as there was a luxurious stand of fennel plants (bushes, actually) on the Burke-Gilman trail across from the J-wing. In a sad turn of events, the plants got cut down a couple of months ago, right around the time Don was hospitalized.

I will miss Don's presence.

I think that perhaps the thing I remember most about Don, aside from how quiet and private he was, was his great patience with the activities of the many scientifically "young" people struggling to learn, hustling and bustling about outside the closed door to his quiet lab. I think the thing I remember most of all, and in fact that I commented on recently to someone I am mentoring, was an incident in which I made a mistake that affected one of Don's experiments. When I went to Don to explain that the error was mine and to apologize, his words were few, spoken quietly. The words themselves are long gone, but his quiet, non-judgmental patience in answering me has stayed with me to this day 30 years later, along with the specific lesson learned, and helped form how I react to the errors people make as they learn science in my lab.

The lesson I got from Don is how to better make your point with a couple of well-chosen words, quietly spoken, than with long lectures, yelling, or derogatory remarks. It was quite notable because I am sure he must have been very unhappy with me at the time. I actually think that he and I both learned something that day, but the most important lesson I carried away was not a lesson about the lab but rather a lesson about people and communication. I think I would be perhaps not as good a mentor if I had not had examples like Don's to counterbalance some of the rather more negative or aggressive approaches to mentoring that I have witnessed along the way. I suspect you will find that many who perhaps did not even know him well will have been touched by him in ways that still affect them many years later.

I am sorry to hear that Don is gone. I hope that his last trip to Paris was truly wonderful. If I were going to pick a last place on Earth to get to go visit, Paris would be high on my list.

I am glad to celebrate with you the memory of Don. I include a few lines, in French, that he would have read with probably a short comment that is hard to guess (may be would have filled in the name of the hotel?).

Don parlait peu, mais savait écouter. Et il avait une mémoire d'éléphant. Il venait quelques fois dîner à la maison. Un soir, il apporte en cadeau une pièce ancienne de l'empire romain, une sesterce d'Antonin le Pieux : "Je sais que tu collectionnes les pièces étrangères d'environ 1$. Ajoutecelle-ci à ta collection". Je l'ai gardée fidèlement, avec le souvenir de Don. Don connaissait le sud de la France, comme sa poche. Il voyageait en train, en bus, et à pied dans un rayon de 10 km autour d'un hôtel. Je lui demandais parfois conseil sur les hôtels, les bons restaurants de l'endroit, et même les horaires de bus. Un jour je l'ai pris en défaut: l'hôtel qu'il recommandait n'existait pas! Deux mois après, il me contacte depuis Seattle. Il venait d'apprendre (comment?) que l'hôtel avait brûlé. Don parlait peu mais n'aimait pas laisser les questions sans réponse.

André et Pierrette Sentenac

Rough translation:

Don spoke little, but knew how to listen. And he had the memory of an elephant. Sometimes he came to dinner at my home. One evening, he brought as a gift an ancient coin for the Roman Empire, a Sesterce of Antonin le Pieux: "I know that you collect foreign coins that have a value of about $1." I treasured it as a souvenir of Don. Don knew the South of France like his pocket. He traveled by train, by bus and by foot in a radius of 10 km around his hotel. On occasion I asked his advice about hotels, good restaurants, and even the bus schedules for specific places. One day I caught him off guard: the hotel that he recommended to me didn't exist! Two months later he contacted me from Seattle. He had just learned (how?) that the hotel burned down. Don spoke little but didn't like leaving questions without an answer.

I shared a small laboratory with Don for about a year and half in the late fifties. My close interaction with him was certainly an enjoyable and profitable scientific experience. He was incredible at mapping yeast genes by tetrad analysis. One day, while cleaning his lab bench, he complained that there were not enough genetic markers. I asked him what he was cleaning his bench with. He replied Roccal, which is a commercial germicide and fungicide. I replied, "why don't get Roccal resistant markers". He immediately pipetted various dilution of Roccal on nutrient plates, and steaked-out a yeast strain. Subsequently he isolated resistant mutants, assigned the mutants to complementation groups, and mapped several of the genes, all within a few weeks. Amazing! The only thing he did faster than mapping genes was driving his sports car.

We planned a family vacation in rural France some years ago. I have always been fascinated by the caves in the Dordogne and lamented to Don that one cannot actually see Lascaux anymore. He immediately suggested an alternative, Rouffignac (which is actually nearby in Perigord). We visited Rouffignac and it was fanatastic.....far larger than Lascaux and very out of the way! We got a complete tour on a little train that the kids (and I !) loved, and saw room after room of fabulous art--mammoths galore. It was an unforgettable experience, and we have Don to thank.

Don was an enigmatic person, both in how he appeared to others and in how he seemed to enjoy puzzling others. At our monthly Seattle Area Yeast Meeting he often appeared but rarely said more than a word. I suspect his conversations were internal and unheard by others, for he always had a countenance of deep contemplation. Perhaps he was contemplating the next trick he might play on someone: an often told, but perhaps apocryphal, story is that he used to send yeast strains to others with the requested genetic markers, but in addition other confounding ones, such as translocations, just to see if the (usually much younger) researcher could figure it out.

Learning that Don is no longer among us was a shock. Don was one of the most dedicated scientists I have ever met. I will never forget the first scientific discussion I had with him (thirty years ago), whilst he was visiting Urs Leupold's lab in Bern. I was so impressed, and he was such a nice person (though I'm not sure I understood half of what he meant). One could say that genetics was his life, and in a sense this would be true. But Don was also open to many things, and knew how to enjoy life. When going to a meeting, he would rarely fail to spent some time in the local museum when there was one. We often met there (occasionally whilst escaping from boring sessions) and it had even become a sort of private joke between us. Yeast genetics, of course, owes him such a lot, though many people may not be aware of it. I suppose it will be difficult to see an ade2-101 red colony without remembering him. Don, we'll miss you.

I first met Don Hawthorne at Bob Mortimer's home in Berkeley. He seemed to me to be just a bit cool about my recent entrance into the field of yeast genetics. He liked the tinyness of the field, just the way it was, and seemed to have the idea that too many people were already engaged in carrying out research on Saccharomyces cerevisiae. He warmed appreciably when I told him how much I admired the work he was doing. At that time Don and Bob were writing their paper on centromere-linked gene maps, which was published in 1960.

The next time I saw Don was in Carbondale in 1962 at what we now call the first international meeting of yeast genetics (although Giovanni Magni preferred to think of it as meeting number zero). Sy Fogel and I had arranged the meeting to be at the University of Southern Illinois with Carl and Gertie Lindegren and Maurice Ogur as the hosts. This was a meeting of about 10 people from the U.S., and one each from Canada and Italy to discuss, and hopefully agree upon, the nomenclature of the genes of yeast. Some of the well-known players in the field were not present, notably Herschel Roman, Boris Ephrussi, Piotr Slonimski, and Otto Winge. Over a very rough three-day period, agreement was achieved. Don played his role in demanding that it be one way only, and Giovanni finally brought the whole enterprise together by drawing an inter-lab list on the blackboard along with the various names, and saying, "We shall call this one his1, right!" pointing his finger at one of us, and then saying "Right!" to the next person, and so on until he had gone around the room. Then Giovanni said, "Okay!" Then he said, "We shall call this one his2, right!" Again around the room until all genes were named, and agreed to, and we could go home. The nomenclature was almost completely what Don and Bob had named the genes--Don's quiet demeanor and stubborn approach to the problem solved the matter in the way he wanted. By that time Carl Lindegren had given up trying to get the nomenclature his way, although no one ever expected him to follow the new rules. Except for his Ph.D. thesis on four-strand crossing over in Neurospora, Carl never had followed rules in any area of genetics, so why should he change now?

Don and Bob were very generous in sending stocks to everyone, and whenever I asked for a stock from Don, I always expected the worst. He had a habit of sending the mutant genes in strains with strange properties which could mask the properties of the mutant genes that I had requested. Don's attitude was, "If he is any good at genetics, he'll figure it out". By good fortune, I never had any problems and could recover whatever mutant genes I had requested from him.

I once visited Don at his home in Seattle. As I recall it, he invited Beth Jones and me over to his house. The most vivid memory of that visit was his showing me a drawerful of salary cheques from the University of Washington in the kitchen that he had never bothered to cash, saying he really didn't need the money.

Don and Bob were instrumental in making yeast genetics accessible to everyone. In 1959, Bob Mortimer and John Johnston had invented the method of using snail gut juice to weaken the ascus walls of yeast before dissecting the spores; Don and Bob together mapped every known gene in yeast, and snail juice helped grease the way. Their mapping papers set the stage for Saccharomyces cerevisae to become the most researched eukaryotic micro-organism, and the first free-living organism to have its genome completely sequenced. The DNA sequence of the yeast genome was completed by December 1995 and put on the internet in March 1996.

The 13th International Conference of Yeast Genetics and Molecular Biology, held in Banff, Canada, in 1986 was dedicated to Donald C. Hawthorne, Robert K. Mortimer, and Seymour Fogel.

My strongest memory of Don is from his attendance at the Seattle Area Yeast Club and departmental seminars. He attended religiously, always sat near the front, and promptly fell asleep with his chin on his chest. It always struck me as profoundly interesting that he made the effort to go to these public meetings, even when he did not have passionate personal interest in them. His presence seemed to say to me that these events were important and that he made the effort to be a part of this community.